A revision of Afrotropical Chyromyidae (excluding Gymnochiromyia Hendel) (Diptera: Schizophora), with the recognition of two subfamilies and the description of new genera.
(Identification and classification)
|Author:||Ebejer, Martin J.|
|Publication:||Name: African Invertebrates Publisher: The Council of Natal Museum Audience: Academic Format: Magazine/Journal Subject: Zoology and wildlife conservation Copyright: COPYRIGHT 2009 The Council of Natal Museum ISSN: 1681-5556|
|Issue:||Date: Dec, 2009 Source Volume: 50 Source Issue: 2|
|Topic:||Event Code: 690 Goods & services distribution; 310 Science & research Advertising Code: 59 Channels of Distribution Computer Subject: Company distribution practices|
|Geographic:||Geographic Scope: United Kingdom Geographic Code: 4EUUK United Kingdom|
Aphaniosoma gaiasicum sp. n. Fig. 45
Etymology: The species name is derived from Gai-As, the type locality.
Diagnosis: Yellow species with 2 pairs of setulae on fr longer than others, dark spot on meron, pale brown stripes on scutum, tarsomeres of all legs yellow and shiny brown spot on ep contrasting with otherwise entirely yellow abdomen. Male postabdomen with distinctive long narrow lobes of distiph and flattened upturned process on basiph.
Head: Yellow, including oc triangle only ocelli ringed black, fr broad but converging anteriorly so that at level of antennae it is about 0.7 as wide as at level of anterior oc, face small, depressed and with distinct, narrow but low facial carina, eye oval lying oblique, gena yellow, at middle about 0.6 height of eye and having several yellow setulae, 2 short setae at vibrissal corner, mouthparts yellow, 4 setulae on fr with 1 pair of longer setulae above lunule and another in front of oc triangle, 3 orb with 1 short setula in front, 1 vti and 1 vte, pvt distinct and convergent, ocellars long, divergent, antenna yellow, most of second segment of arista dark.
Thorax: Scutum and scut with yellow ground but with pale brown longitudinal stripes on scutum, mtn brown, pleura all yellow except for a pale brown inverted triangle on lower katepisternum and dark brown spot on meron. Chaetotaxy: 1 pprn, 1 posthu, 1 ihu, 2 ntpl, 0+4 ia, 2+4 acrs, 3+4 dc only the posterior one well-developed, prpl not seen, 1 pa, 2 pairs of marginal scut, 1 katepisternal, 1 anepisternal, all setae pale yellow. Wing: Hyaline and uniformly microtrichose, veins pale yellow, distance between [R.sub.2+3] to [R.sub.4+5] on costal margin about 0.4 that between [R.sub.4+5] and [M.sub.1+2], distance between crossveins about 1.2x length of posterior crossvein and length of apical section of Cu about 2.8x length of posterior crossvein. Haltere yellow.
[FIGURE 45 OMITTED]
Legs: Yellow and fine yellow setulose, tarsomere 5 of all legs yellow.
Abdomen: Yellow with narrow, brown anterior margins to all tg. Postabdomen: Yellow with shining brown ep spot lateral to cerc.
As male. (Postabdomen not dissected due to poor condition.)
Length: [male] body 1.2 mm, wing 1.4 mm; [female] body 1.4 mm, wing 1.5 mm.
Holotype: [male] NAMIBIA: Khorixas District, Gai-As spring, 20[degrees]46'01"S:14[degrees]01'12"E, 22.x.1998, A.H. Kirk-Spriggs & E. Marais (in glycerine, NMNW).
Paratypes: 2 [female] in alcohol, same data and depository.
Aphaniosoma gallagheri Ebejer, 1996 Fig. 46
Aphaniosoma gallagheri: Ebejer 1996: 292.
The finding of this species in The Gambia and Cape Verde was a surprise, given its apparent absence from intervening countries. It was described from southern Oman. The species is illustrated again using specimens from Cape Verde (male) and The Gambia (female) in order to supplement the original figures with more detail. The female is very similar to the male in external features. Postabdomen: tg 7 divided into lateral plates with relatively long setulae; tg 8 in two small sclerotized lateral plates, st 7 membraneous; hypr small barely discernable except for minute setulae.
Material examined: CAPE VERDE: 31 [male] 11 [female] Sal, Saline Pedra Lumen, 5-7.i.1985, P. Ohm (in alcohol, FBUB); 1 [female] Santiago, b. Variante, "Trockenribeiro und Staatsfarm", 2.xi.1982, P. Ohm (in alcohol, FBUB). THE GAMBIA: 1 [male] 2 [female] Kotu stream, about 3 km SW Bakau, 23.xi.1977, swept in vegetation, Lund University Systematics Department, Cederholm, Danielsson, Hammarstedt, Hedqvist, Samuelsson (MZLU); 1 [female] same data but 23.ii.1977, in vegetation and freshwater (MZLU); 29 Bakau, at Tropic Bungalow, 16-18.xi.1977, swept in meadow rich in flowers at the beach, Lund University Systematics Department, Cederholm, Daniels-son, Hammarstedt, Hedqvist, Samuelsson (MZLU); 1 [female] West Division, Bakau Carmalou Corner, 22.xi.1993, on Phragmites in saltmarsh, J.C. Deeming (NMWC); 1 [female] West Division, 26.xi.1993, on grasses bordering Kotu stream, J.C. Deeming (NMWC); 1 [male] West Division, Lamin Bridge, Lamin Bolan, 2.viii.1997, D.J. Mann (NMWC).
[FIGURE 46 OMITTED]
Distribution: Cape Verde, The Gambia, Oman (Dhofar).
Aphaniosoma harteni: Ebejer 1996: 293.
Described from Yemen and later found in Tunisia, this species is unique in the Afrotropical fauna. It is the only representative of the group of species with a strong presutural dc seta. This species group is distributed across the Mediterranean, with four species recorded from the Canary Islands and one from Mongolia.
Distribution: Tunisia, Yemen.
Etymology: Named after Ashley Kirk-Spriggs, who collected many Chyromyidae in Namibia.
Diagnosis: Yellow species with black ringed ocelli, elongate slender eyes, dark longitudinal scutal stripes, scut with dark lateral margin reaching base of subapical setae; abdomen yellow with light brown bands dorsally on tg; male postabdomen with large shiny dark brown phallic structures easily visible, owing to their contrast with yellow tg and st. Male postabdomen with setulose surs and prg, and bilobed distiph.
Head: fr, oc triangle and ocp entirely yellow, only ocelli black; eye narrow and horizontal, about 3 times as long as high; gena yellow and pale yellowish white setulose, about equal to height of eye; face very small and membranous with shallow linear carina; antenna including first segment of arista yellow, second aristal segment dark brown; mouthparts yellow. Chaetotaxy: 3 reclinate orb with short setula in front, 2 long ocellars widely spaced almost parallel, 10 fr setulae and a pair of long setae in front of oc triangle; 1 vti and 1 vte, pvt convergent.
[FIGURE 47 OMITTED]
Thorax: Scutum yellow and pale yellow pollinose with dark grey scutal stripes confluent anterior to transverse suture; scut yellow with brown lateral margin up to subapical setae; mtn dark and pleura yellow with dark greyish brown triangular mark on katepisternum and spot on meron. Chaetotaxy: 2+5 dc only hindmost developed to a seta, 0+4 acrs in two irregular rows, 1 pprn, 1 ihu, 1 posthu, 1 ia, 2 ntpl, 1 pa, 1 anepisternal and 1 katepisternal with a few setulae near each.
Wing: Hyaline with yellow veins becoming darker towards apex; distance between [R.sub.2+3] and [R.sub.4+5] about 0.5 that between [R.sub.4+5] and [M.sub.1+2]; distance between crossveins equal to length of posterior crossvein. Haltere pale yellow.
Legs: No special modifications, yellow and yellow setulose, tarsomere 5 brown.
Abdomen: tg pale brown with yellow hind margins which occupy about 1/4 length of tg dorsally, but become broader on sides.
Postabdomen: cerc small yellow and pointed; distiph large shiny and dark.
Very similar to male; eye less elongated. (Postabdomen not dissected.)
Length: [male] body 1.1 mm, wing 1.0 mm; [female] body 1.2 mm, wing 1.3 mm.
Variation: The male paratype has the eye only twice as long as high and the tarsomeres being all yellow.
Holotype: [male] NAMIBIA: Luderitz District, Angra Bay, 26[degrees]37'11"S:15[degrees]11'42"E, sweeping lush vegetation at effluent run-off stream, 27.iii.1998, A. Kirk-Spriggs (NMNW). Paratypes: NAMIBIA: 4 [male] 2 [female] same data as holotype (NMNW); 1 [male] 1 [female] Otjitambi farm, 47 km ESE Kamanjab, 13-15.ii.1972, Southern African Expedition (W37) BM 1972-1 (BMNH). SOUTH AFRICA: Western Cape: 1 [male] (headless) 1 [female] Sandfontein Coast, west of Van Rhynsdorp, 15-17.x.1964, B. & P. Stuckenberg (NMSA).
Aphaniosoma lamellatum Collin, 1949
Aphaniosoma lamellatum: Collin 1949: 137.
This is one of the species that Collin described from Egypt. Although it has not been found on mainland Africa south of the Sahara, it has been recorded from Oman (Ebejer 1996) and is included here for completeness as it could easily be more widely distributed. The long yellow lamellae referred to by Collin are the surs, and these are indeed long and broad with somewhat sinuous margins as illustrated by him.
Distribution: Egypt, Oman.
Aphaniosoma micropygum sp. n. Figs 48, 49
Etymology: From Greek micros (small) and Latin pyga (buttocks), and refers to the particularly small hypopygium.
Diagnosis: Yellow species with brown longitudinal stripes on scutum and short pra setula; brown tarsomeres 5; abdomen yellow with tg having brown band along anterior margin, male postabdomen particularly small, yellow and with most structures translucent; lower margin of tg 6 with posterior indentation.
Head: Entirely yellow, only ocelli ringed with black; antenna yellow, arista missing, mouthparts yellow; face very small and sunken; gena yellow and pale setulose; gena about 0.6 height of eye, which is long oval and lies almost horizontal. Chaetotaxy: 2 orb and 2 setulae in front of these, 8 scattered short fr setulae and 1 pair of long setae in front of oc triangle; 1 vti and 1 vte and short pvt; postoculars in 1 row.
Thorax: Yellow, with brown longitudinal stripes on scutum, scut yellow, mtn brown, pleura yellow except for brown triangular patch on katepisternum, narrow brown line on anepisternum and anepimeron near suture, meron all brown. Chaetotaxy: 1 pprn, 1 posthu, 1 ihu, 3+4 dc only hindmost developed to a seta, 0+4 acrs, 1+3 ia setulae, 1 pa, 1 anepisternal and 1 katepisternal, latter with vertical row of 4 short setulae down middle of sclerite.
Wing: Hyaline and uniformly microtrichose, veins pale brown to yellow, distance between [R.sub.2+3] to [R.sub.4+5] on costal margin about 0.7 that between [R.sub.4+5] and [M.sub.1+2], distance between crossveins about 1.2x length of posterior crossvein and length of apical section of Cu about 2.8x length of posterior crossvein. Haltere pale yellow.
Legs: All yellow and yellow setulose, tarsomere 5 brown.
Abdomen: Yellow, with brown bands along anterior half of tg and pale hind margin becoming broader at sides.
Postabdomen: Very small and pale hypopygium; most structures translucent; hyp particularly small and narrow; surs and prg run parallel to each other, surs originating from lower end of ep and prg from posterior end of hyp where this fuses with ep; ep itself small, trapezoidal and invaginated into space within tg 6; pregenital st small and narrow forming loop around ventral hypopygial opening, posteriorly at midline forms a keel in front of distiph.
[FIGURE 48 OMITTED]
[FIGURE 49 OMITTED]
Very similar to male. Postabdomen: tg 7 broad with long setulae; tg and st 8 sclerotized and divided, latter of characterisitc shape, st 7 elongate oval with distinct setulae; s with thinly sclerotized first part of duct; cerc small.
Length: [male] and [female] body 1.4 mm, wing 1.5 mm.
Holotype: [male] NAMIBIA: Swakopmund [22[degrees]40'S:14[degrees]33'E], general sweeping, 26-30.i.1972, Southern African Expedition (W25) B.M. 1972-1 (BMNH).
Paratypes: 1 [male] 7 [female] same data as holotype (BMNH).
Aphaniosoma pullum sp. n. Figs 50, 51
Etymology: From Latin pullum (blackish), and refers to this species being predominantly dark coloured.
Diagnosis: Extensively greyish black species with brown third antennal segment and tarsomeres of all legs yellow in [male] but often brown in [female], scut setae longer than length of scut, ep shiny dark brown contrasting with the pale cerc; surs unusual, with very short lobes and bilobed distiph projecting backwards rather than forwards.
Head: Yellow (rather dusky due to postmortem changes), ocp black, oc triangle black, fr broad but markedly converging anteriorly so that at level of antennae it is about 0.5 times as wide as at level of anterior oc, face small, depressed and with distinct, narrow but low facial carina, eye oval lying oblique, gena yellow, narrow, at middle about 0.4 height of eye and having numerous pale yellow setulae, 2 short setae at vibrissal corner, mouthparts yellow, 4 setulae on fr with 1 pair of longer setulae in front of oc triangle, 4 short orb getting progressively shorter from back to front, 1 vti and 1 vte, pvt distinct and convergent, ocellars parallel, antenna brown (though not as dark chocolate brown as in atriceps), arista dark, only extreme base of segment 1 yellow.
Thorax: Scutum dark grey, scut dark brown, mtn entirely dark grey, pprn and ntpl yellow, pleura mostly dark grey, but for narrow yellow margins to sutures. Chaetotaxy: 1 pprn, 1 posthu, 1 ihu, 2 ntpl, 0+2-3 ia, 1+5 acrs, 4+5 dc only the posterior one well-developed, prpl minute, 1 pa, 2 pairs of marginal scut, all longer than length of scut, 1 katepisternal with shorter seta in front and several setulae on anterior part of sclerite, 1 anepisternal, all setae pale at front of body becoming darker brown on posterior parts.
[Figure 50 Omitted]
Wing: Hyaline and uniformly microtrichose, veins brown, distance between [R.sub.2+3] to [R.sub.4+5] on costal margin about 0.6 that between [R.sub.4+5] and [M.sub.1+2], distance between crossveins about 1.3x length of posterior crossvein and length of apical section of Cu about 2.8x length of posterior crossvein. Haltere yellow.
[FIGURE 51 OMITTED]
Legs: Yellow and fine yellow setulose, tarsomere 5 of all legs yellow.
Abdomen: Dark brown with narrow, pale yellow hind margins to all tg. Postabdomen: Yellow with upper part of ep shiny brown on each side of cerc.
As male, but several specimens have tarsomeres 5 darkened. Postabdomen: tg 7 divided, st 7 elongated oval with distinct setulae; cerc appear fused to tg 8; hypr distinct, but no special features; st 8 in two small lobes.
Length: [male] body 1.0 mm, wing 1.1 mm; [female] body 1.0 mm, wing 1.2 mm.
Holotype: [male] NAMIBIA: Luderitz [26[degrees]39'S:15[degrees]10'E], alt. 20 m, 14.ix.2003, A. Freidberg (TAUI). Paratypes: 1 [male] 5 [female] same data as holotype (TAUI).
Aphaniosoma suboculicauda Frey, 1958 Fig. 52
Aphaniosoma suboculicauda: Frey 1958: 34.
Frey's description, though accurate on most points, has some errors and his illustration of this species is difficult to interpret. Given that so many species have been described since that time, an examination of the types was mandatory. The series of specimens that I received for study from MZH consisted of 11 specimens, none of which is labelled by Frey with the name of the species. The two specimens with a species's name are those that I examined and labelled in 1997 (Ebejer 1998) for the revision of the Palaearctic species of this genus. Although not labelled by Frey as types, all the specimens I examined bear the same data as given in his paper for an unspecified number of specimens. There is no other species of Aphaniosoma that he refers to. For this reason, I consider them all to constitute the type series as Frey implied in his article.
The holotype [male] carries the following labels: "Holotypus" [red label]; "spec.typ." [typed], "holotyp [sic]" [hand written below]; "Ins. Cabo Verde, Maio Rib. da Lagoa, 2.2.1954, Lindberg". It is indeed a male in perfect condition with the phallic complex partly extruded. The apex of the distiph is not visible, but no black structures can be made out, even though the postabdomen is pale and semi translucent. In such cases, dark internal structures are visible through the integument, although detail might not be appreciated. The basiph, surs and psg are pale throughout. The prg cannot be discerned. Frey describes a pair of black spots on tg 6. None of the males in this series has a black spot. The small brown spot that is present is on the ep. Frey's illustration of the male postabdomen and his comments upon it raise the possibility that the specimen it comes from might not be the same species as the one designated as the holotype. However, since all the type series represents one species, I am more inclined to believe that Frey rather overstated the features and in his illustration of the distiph over-inked the dark areas. There are no hairs on the distiph either in the male I dissected or indeed in any species of Aphaniosoma that I have seen with the sole exception of trilobatum sp. n. (see below). Dr Vilkamaa informs me that he has been unable to trace the slide preparations of the two abdomens that are missing from the type series and upon which Frey based his illustration. For these reasons, I give an illustration of a male paratype that I dissected in order to allow this species to be properly recognized. I labelled the remaining specimens as paratypes since they are so referred to, albeit indirectly, in Frey's paper.
[male] "Ins. Cabo Verde, Maio Rib. da Lagoa, 2.2.1954, Lindberg", in very good condition.
"Paratypus" [red label]; "[male]" [typed in large font]; "Ins. Cabo Verde, Boavista, Fundo de Figueiras, 30.1.1954, Panelius". This specimen is in very good condition, but the abdomen was neatly removed from tg 3 onwards and therefore I cannot confirm the sex. The colouration of the head and thorax, the proportions of the wing veins and spaces between them and the chaetotaxy suggest that this is the same species as the holotype.
"Paratypus" [red label]; "pinz" [hand written, deciphered correctly?]; "Ins. Cabo Verde, Maio Pedro Vas, 3.2.1954, Lindberg". This specimen is greasy and very badly damaged: headless, all right legs missing, tarsomeres of middle and hind leg missing on left side, right wing folded, left wing mostly missing and apex of abdomen missing (apparently through damage). There is nothing about the remains of this specimen that suggests it is a different species to the above. There are three more males bearing the same data and all are the same species as this. I dissected one of these.
[FIGURE 52 OMITTED]
"Paratypus" [red label]; "18" [typed in large font]; "Ins. Cabo Verde, Boavista Rabil, 31.1.1954, Lindberg". The specimen is damaged: headless and left wing folded. The abdomen has been cleanly removed from tg 3 onwards. Therefore, I cannot determine the sex. This is the same species as the rest.
"Paratype, [female] Aphaniosoma suboculicauda Frey, det. M.J. Ebejer, 1997"; "Ins. Cabo Verde, Boavista, Fundo de Figueiras, 30.1.1954, Panelius". Apart from missing the left wing the specimen is in good condition.
"Paratype, [female] Aphaniosoma suboculicauda Frey, det. M.J. Ebejer, 1997"; "Ins. Cabo Verde, Maio Pedro Vas, 3.2.1954, Lindberg". The specimen has the scutum badly cracked, but the parts are still present, the head is shrivelled, but the chaetotaxy can be made out. This is a darker specimen than all the foregoing. It has more extensive darkening of pleura, scutal stripes and abdominal tg. The chaetotaxy is very similar to the foregoing specimens and I consider the chromatic differences of minor importance and well within the variation commonly seen in species of this genus.
Finally, a [male] also belonging to the same species as above, bears the following label: "Ins. Cabo Verde, S. Tiago Achada, Robao Areia, 11.2.1954, Lindberg". It has a rather shrivelled head and the scutum is broken, but all parts are present.
Distribution: Cape Verde.
Aphaniosoma trilobatum sp. n. Figs 53, 54
Etymology: From Latin tri- (three-) and lobatus (lobed), and refers to the shape of the male postabdominal processes.
Diagnosis: Predominantly yellow species with 6 dark grey longitudinal stripes on scutum, rather longer setae and setulae, including those on gena, than in congeners, fr with 2 pairs of longer setulae, tarsomere 5 of all legs brown. Male postabdomen with three broad yellow lobes, easily visible without dissection and appearing to be part of same structure. However, when dissected these are surs, prg and tip of ep.
Head: Yellow, ocp dark brown except for broad yellow postocular margin and wedgeshaped area from vertex to neck, oc triangle black, fr broad but converging anteriorly so that at level of antennae it is about 0.7 as wide as at level of anterior oc, face small, depressed and with distinct, narrow but low facial carina, eye oval lying oblique, gena yellow, at middle about 0.5 height of eye and having numerous rather longer yellow setulae, 2 short setae at vibrissal corner, mouthparts yellow, 8 setulae on fr with 1 pair of longer setulae above lunule and another in front of oc triangle, 3 orb with 1 short setula in front, 1 vti and 1 vte, pvt distinct and convergent, ocellars long, divergent, antenna yellow, most of second segment of arista dark.
[FIGURE 53 OMITTED]
Thorax: Scutum and scut with yellow ground but with 6 dark grey longitudinal stripes on scutum, middle pair confluent anteriorly, scut a little pale brownish on disk, mtn brown, pprn and ntpl yellow, pleura mostly yellow except for a brown patch on anterior and lower part of anepisternum and an inverted dark grey triangle on each lower part of katepisternum and meron, base of haltere black. Chaetotaxy: 1 pprn, 2 posthu, 1 strong ihu, 2 ntpl, 0+4 ia, 2+5 acrs, 3+4 dc only the posterior one well-developed, prpl short, 1 pa, 2 pairs of marginal scut, 1 katepisternal with shorter seta in front and several setulae on anterior part of sclerite, 1 anepisternal, all setae pale brown and rather longer than those in other species described in this article.
Wing: Hyaline and uniformly microtrichose, veins pale brown, distance between [R.sub.2+3] to [R.sub.4+5] on costal margin about 0.4 that between [R.sub.4+5] and [M.sub.1+2], distance between crossveins about 1.3x length of posterior crossvein and length of apical section of Cu about 2.3x length of posterior crossvein. Haltere yellow.
Legs: Yellow and fine yellow setulose, tarsomere 5 of all legs brown.
Abdomen: Brown with narrow, pale yellow hind margins to all tg.
Postabdomen: Yellow with brown ep spot above and lateral to cerc and with distinctly visible large yellow trilobed surs.
As male, but without secondary sexual characters. Postabdomen: tg 7 divided, st 7 elongate oval with setulae only at apex; tg 8 and st 8 each divided into small sclerites laterally; hypr not identified.
Length: [male] body 1.1 mm, wing 1.2 mm; [female] body 1.5 mm, wing 1.6 mm.
Holotype: [male] NAMIBIA: Walvis Bay, alt. 5 m, 22[degrees]60'S:14[degrees]28'E, 6.ix.2003, A. Freidberg (TAUI). Paratypes: NAMIBIA: 5 [male] 7 [female] same data as holotype (TAUI); 1 [male] 1 [female] Luderitz, alt. 20 m, 14.ix.2003, A. Freidberg (TAUI); 1 [female] Gobabeb, Kuseib R., 23[degrees]34'S:15[degrees]03'E, 5.ix.2003, attacked by Laboulbeniales, A. Freidberg (NMWC); SOUTH AFRICA: Western Cape: 1 [male] De Hoop Nature Reserve, alt. 0-200 m, 34[degrees]27'S: 20[degrees]25'E, 10-13.x.1994, R. Danielsson (MZLU).
[FIGURE 54 OMITTED]
Aphaniosoma ugabensis sp. n. Fig. 55
Etymology: The species name is derived from the type locality, Ugab River in Namibia. Diagnosis: Almost entirely yellow species with 2 pairs of long setulae on fr, only dark areas are small rectangular patch at anterior end of scutum just above occipital foramen and on metanotum. There are no external features to help in the identification of this species. Male postabdomen with long narrow pointed psg and short truncate prg; surs reduced to an inconspicuous lobe; pregential st with distinct midline ventral projection.
Head: Entirely yellow, only ocelli ringed with black, fr broad, converging anteriorly so that at level of antennae it is about 0.7 as wide as at level of anterior oc, face small, depressed (carinal area not visible), eye narrow, oval, lying oblique, gena yellow, shrivelled, probably about half height of eye and having numerous yellow setulae, setae at vibrissal corner not seen, mouthparts yellow, 4 setulae on fr with 2 pairs of longer setulae, one above lunule and one in front of oc triangle, 2 short orb and in front of these 2 very short fine setulae,1 vti and 1 vte,pvt distinct and convergent, oc long, divergent, antennal segment 3 yellow, arista dark brown on apical 2/3 of segment 2 of arista.
Thorax: Scutum and scut yellow with pale brown longitudinal stripes on posterior half of scutum and brown patch on extreme anterior scutum above neck, mtn dark brown, pleura yellow. Chaetotaxy: 1 pprn, 1 posthu, 1 ihu, 2 ntpl, 0+3 ia, 2+3 acrs, 3+4 dc only the posterior one well-developed, prpl not seen, 1 pa, 2 pairs of marginal scut, 1 katepisternal, 1 anepisternal, all setae pale, with shorter ones and setulae inconspicuous.
Wing: Hyaline and uniformly microtrichose, veins yellow, distance between [R.sub.2+3] to [R.sub.4+5] on costal margin about 0.5 that between [R.sub.4+5] and [M.sub.1+2], distance between crossveins about equal to length of posterior crossvein and length of apical section of Cu about 3x length of posterior crossvein. Haltere yellow.
[FIGURE 55 OMITTED]
Legs: Yellow and fine yellow setulose, tarsomere 5 of anterior leg brown. Abdomen: Yellow on all tg.
Postabdomen: Yellow with no distinctive external features. Female.
As male, but several specimens have the middle third of femora diffusely infuscated dorsally. (Postabdomen not dissected.)
Length: [male] and [female] body 1.2 mm, wing 1.4 mm.
Holotype: [male] NAMIBIA: Omaruru District, Ugab R., 2 km W Brandberg Wes, 20[degrees]58'05"S:14[degrees]06'36"E, 23.x.1998, sweeping, E. Marais (in glycerine, NMNW). Paratypes: 2 [male] 8 [female] same data as holotypes (in alcohol, NMNW). Remarks. The holotype is not in good condition due to some shrivelling and broken setae.
Genus Krifomyia gen. n.
Type species: Rhicnoessa minutissima Bezzi, 1908: 200, here designated.
Etymology: From Greek krifos (secret, secluded) and myia (fly), and refers to the genus being rare and unobtrusive.
Diagnosis: Well-developed, broad facial carina extending from lunule to just below antennae where it broadens laterally leaving antennae in deep pits; head short with rather inflated fr that has parallel margins and which, in profile, is visible all along eye margin from vertex to antenna, gena higher than height of eye which, in profile, is round to oval and small relative to head size, when compared to other genera; pvt present, ihu minute, posthu short, sa and katepisternal setae absent, veins [R.sub.4+5] and [M.sub.1+2] converging slightly towards apex, apicoventral seta on mid tibia present.
Head: Rectangular in profile and almost twice higher than long; fr broad and parallelsided, protruding above and in front of eye margin when seen in profile; finely setulose; gena deep, at middle equal to or greater than height of eye and with wide post genal dilation; fine pale setulose, postgenal seta absent; face long with very broad carina extending from lunule to just below antennae, where it widens; thus, antennae lie in deep fossae, with segments 1 and 2 very short, segment 3 rounded except for shallow dorsal excavation near insertion of arista; arista with 3 segments and shorter than usual for the family; 2 short reclinate orb, 1 vti and 1 vte, both shorter than usual for this family, oc setae divergent, pvt minute and parallel, sometimes absent, few postocular setulae in one row; mouthparts all yellow, well-developed with long proboscis, geniculate labella and narrow elongated palp.
Thorax: A little shorter and broader than usual for the family. Chaetotaxy: 1 short pprn and 1 short posthu, a minute but distinctly incurved ihu, 2 ntpl, 1 pa, 1 short post ia, 1 dc, acrs very short and in more than 2 irregular rows, 1 anepisternal closer to upper margin; absent sa and katepisternal setae; scut with 1 pair basals and 1 pair subapicals, all about as long as dc.
Wing: Hyaline, veins very pale yellow, only Cu along discal vein a little darkened; RM stem vein thickened; anal cell pointed apically along anal vein; costal setulae all fine and pale; [R.sub.2+3] and [R.sub.4+5] very slightly divergent. Haltere yellow with creamy white knob. Legs: Entirely dull yellow with short pale setulae, only claws darkened; apicoventral seta of mid tibia present and longer than diameter of tibia at apex; legs without dilated segments or special modifications.
Abdomen: All tg normally sclerotized and with a few very fine setulae scattered thinly on all segments, but on tg 6, which is as long as tg 5, stronger and more numerous setulae present.
Postabdomen: Male hypopygium very small with reduced appendages, short ph apd and markedly simplified ph compared to the rest of the family. Female abdomen with 8 segments, sclerotized st 8; 2 unpigmented s.
Remarks. This genus belongs to the Aphaniosominae because of the following combination of characters: absent incurved lower orb, concave ocp, long apicoventral seta on mid tibia, anepisternal seta above middle of posterior margin, incurved ihu setula, absent hemispherical ep. With Paraphaniosoma it shares the post ia seta and more than two rows of scutal setulae between dc lines. However, it differs in the much simplified hypopygial structures, absent sa and katepisternal setae, markedly different shaped head, well-developed proboscis and in the wing having the costal vein with spinescent setulae and a thickened hu crossvein.
Distribution: Atlantic coast of southern Africa (Namibia) (Fig. 70).
Ecology: Krifomyia, known from a single species so far, is rare and by all accounts a rather cryptic inhabitant of hot dry coastal regions.
Krifomyia minutissima (Bezzi, 1908), comb. n. Figs 56, 57
Rhicnoessa minutissima: Bezzi 1908: 200.
The type material consists of 1 [male] and 1 [female] from Namibia, both in poor condition, preserved together in 70% alcohol. The male was partially dissected by Munari (Munari 1994) who was the first to recognize that this species did not belong to the Tethinidae (now a subfamily of the Canacidae), but to the Chyromyidae. The head of the male was separated from the body, as were a wing (the other missing) and the abdomen. The thorax has one of each of the fore, middle and hind legs, the rest are missing. The abdomen is squashed and partly macerated. The female is badly shrivelled, but retains most of its parts. Nevertheless, there are sufficient discernible characters to allow me to confidently assign to this species a number of specimens collected more recently, also from Namibia, which are better preserved. The descriptions and drawings are based on these specimens.
Head: All yellow; in profile rectangular and almost twice higher than long; fr broad and parallel-sided, about 3x width of eye viewed from in front, protruding above and in front of eye margin viewed in profile; with about 10 short and fine setulae scattered on it; eye oval, gena deep, at middle equal to or greater than height of eye and with wide postgenal dilation; several short fine pale setulae scattered over lower part, postgenal seta absent; face with very broad carina extending from lunule to below antennae, leaving from this level to buccal margin a relatively large rectangular sclerotized area; antenna in deep fossa, segments 1 and 2 not visible, segment 3 rounded except for shallow dorsal excavation near insertion of arista; arista with 3 segments, a little shorter than is usual for this family, first 2 segments and base of segment 3 pale; 2 short reclinate orb and one fine setula in front, 1 vti and 1 vte, both shorter than is usual in this family, oc setae divergent, pvt minute and parallel, sometimes absent, few postocular setulae in one row; mouthparts all yellow, well-developed with long proboscis, geniculate labella and narrow elongated palp.
Thorax: Yellow, often with longitudinal pale brown scutal stripes, pleura always yellow. Chaetotaxy: scut yellow dorsally but with dark spot laterally at base continuous with a dark brown line along entire lower border of scut; 1 short pprn and 1 short posthu, a minute but distinctly incurved ihu, 2 ntpl, 1 pa, 1 short post ia, 1 dc, scutal setulae very short and fine in 4-6 irregular rows, 1 anepisternal closer to upper margin; absent sa and katepisternal setae; scut with 2 pairs of marginals, basals and subapicals all about as long as dc.
[FIGURE 56 OMITTED]
[FIGURE 57 OMITTED]
Wing: Hyaline; veins very pale yellow, only Cu along discal vein a little darkened; RM stem vein thickened, anal cell pointed apically along anal vein; costal setulae all fine and pale. Haltere yellow with creamy white knob.
Legs: Entirely dull yellow with short pale setulae, only claws darkened; apicoventral seta of mid tibia present and longer than diameter of tibia at apex; legs without dilated segments or special modifications.
Abdomen: All tg with brown dorsal bands, leaving hind and lateral margins pale yellowish white; a few very fine setulae scattered thinly on all segments, but segment 6, which is as long as segment 5, has stronger and more numerous setulae.
Postabdomen: Very small and poorly sclerotized except for margins of hyp; ep poorly developed, but complete dorsally; cerc minute, translucent and very finely setulose; surs elongate, fused with ep and finely setulose;ph sclerotized but simple, almost tubular; ph apd very small and indistinct, only anterior apex sclerotized, gonites and ej apd could not be identified.
Very similar to male. All abdominal segments with midline sclerites. Postabdomen: tg 7 complete; st 8 sclerotized with small posteromedial lobe and setae ventrally; hypr small oval with fine setulae apically; cerc in proportion to adjacent structures rather large and setose apically.
Length: [male] and [female] body 1.3 mm, wing 1.2 mm.
Variation: Presence and colour of scutal stripes variable, number of setulae on fr and acrs a little variable, otherwise all specimens examined are remarkably similar.
Type material examined: syntypes NAMIBIA: [male] "Klein Namaland, SW Africa, leg. Shultz (Humbolt)" (ZMHB); 1 [female] same data (ZMHB).
Other material examined: NAMIBIA: 1 [female] Luderitz District, Agate Beach, 10 km N Luderitz, alt. 3 m, low coastal vegetated dunes, 18.ii.1974, M.E. & B.J. Irwin (NMSA); 69 Skeleton Coast Park, Unjab R. Delta, 20[degrees] 11'S:13[degrees] 13'E, 18-26.vii.1990, C.S. Roberts, pitfall traps (NMWC); 1 [male] Luderitz District, [missing locality] 27[degrees]34'S:15[degrees]28'E, 15-25.xi.1993, E. Marais, pitfall traps (in glycerine, NMWC); 1 [male] 3 [female] same data (in alcohol, NMNW); 1 9 Walvis Bay, alt. 5 m, 22[degrees]60'S:14[degrees]28'E, 6.ix.2003, A. Freidberg (TAUI).
Genus Paraphaniosoma gen. n.
Type species: Aphaniosoma sexvittata Lamb, 1914, here designated.
Etymology: From Latin par (like) and Aphaniosoma, the genus which it resembles.
Diagnosis: Separation of genus from Aphaniosoma is easy on basis of well-developed pra and post ia setae, scutal setulae in six or more rows between dc lines, only one strong dc and reduced or absent pvt. As in Aphaniosoma, strong apicoventral seta present on mid tibia, longer than diameter of tibia at apex.
Head: Margins of fr converge anteriorly; ocp concave, gena much narrower than height of eye; 1-5 orb reducing in size from back to front, all reclinate; 1 strong vti and 1 strong vte, pvt absent or minute and inconsistently present within a species; mouthparts normal; antenna as for the rest of the family, but arista 2-segmented.
Thorax: Appears somewhat elongated, a little less arched and usually with distinct longitudinal stripes on scutum. Chaetotaxy: 1 to 2 pprn, 1 ihu, 1 posthu, 2 ntpl, 1 pra, 1 sa, 1 pa, 1 post ia, 0+1-2 dc, scutal setulae in 4-8 rows with long prscut, 1 anepisternal at top of posterior border, 1 katepisternal at upper hind corner, no prpl; scut with basal and subapical pair.
Wing: Hyaline; crossveins and vein separating discal from basal cell translucent; [R.sub.4+5] and [M.sub.1+2] parallel or nearly so. Haltere without special modifications.
Legs: Normal, with no special modifications, mid tibia ventrally at apex with strong seta.
Abdomen: 6 visible tg in male, 7 or 8 in female; in both sexes, tg more or less uniformly short setulose.
Male postabdomen of the Aphaniosoma s. str. type, i.e., a small, partly internalized ep and markedly modified gonites and pregenital st. Female postabdomen with large tg 7, cerc separated from segment 8, which has a pair of sclerotized st plates.
Distribution: East Palaearctic, Afrotropical (Fig. 70), Oriental, ?Australasian. Most species are undescribed.
Ecology: Little is known about this genus, but the few records for Paraphaniosoma seem to be mainly from coastal and estuarine localities in hot tropical and subtropical climates. As more species undoubtedly will be discovered, this postulation could change.
Paraphaniosoma sexvittatum (Lamb, 1914), comb. n. Figs 58, 59
Aphaniosoma sexvittata: Lamb 1914: 356.
The original description of P. sexvittatum, although accurate, lacks detail and would not satisfactorily differentiate this species from other undescribed species I have seen from the East Palaearctic and Oriental regions (Papua New Guinea, Malaysia, Taiwan). Therefore, a more detailed redescription is provided with figures of the male and female postabdomen.
The detailed description of the characters given by Lamb (1914), and the characters given in the key to genera provided in this article, serve to identify this species with relative ease. Additional characters and illustrations are given below, based on more material of this species from Aldabra (Seychelles).
Head: Yellow, oc triangle yellow, but ocelli ringed with black; ocp brown except for broad postocular margin; fr narrow, at level of anterior oc about 0.4 width of head and at level of antennae about 0.6 width of that at level of anterior oc; eye elongate oval, lying oblique. Chaetotaxy: 2 reclinate orb with 3 short pale setulae in front, 1 vti and 1 vte, minute pvt (absent in some specimens); ocellars long and divergent with pair of short setulae behind these in between posterior ocelli; 14 setulae scattered on fr, but 6 of these in front of oc triangle about as long and strong as anterior orb; postoculars in one row; gena pale yellow and very pale yellow setulose with stronger setulae on oral margin, gena at middle about 0.5 highest diameter of eye; mouthparts yellow, palpus narrow; face mostly membranous with a very thin and shallow facial carina; antenna yellow, second segment paler with very thin short and pale dorsal seta; arista all brown, bare.
Thorax: Yellow, paler on pleura, scutum with 6 longitudinal dark brown stripes all separated from each other and from a brown patch at anterior end of scutum; middle stripes run from just anterior to transverse suture to scut, lateral to these and from behind transverse suture, a stripe runs between dc and ia lines of setulae on each side and another between sa and ia rows; a brown tiangular spot lies medial to pprn and confluent or narrowly separated from anterior dark scutal patch; anepisternum with brown streak at its lower margin and anepimeron with brown streak at its anterior margin, mtn brown, scut mostly yellow except for broad pale brown patch on disc and dark brown spot on sides at base. Chaetotaxy: 2 pprn, 2 ntpl, 1 posthu, 1 short ihu, 1 pra, 0+2 dc (anterior one short), 1 short sa, 1 long pa and 1 long post ia, scutal setulae in 4-6 irregular rows between dc lines, with stronger prscut pair, scut with basal and subapical pair, 1 anepisternal at upper hind margin and 1 katepisternal at upper hind corner.
[FIGURE 58 OMITTED]
[FIGURE 59 OMITTED]
Wing: Hyaline with pale brown veins; [R.sub.2+3] somewhat undulating at middle, [R.sub.4+5] and [M.sub.1+2] parallel and on costal margin separated by a distance equal to half distance between [R.sub.4+5] and [M.sub.1+2]; crossveins and vein between discal and basal cell translucent pale yellow, distance between crossveins equal to 1.2x length of posterior crossvein. Haltere yellow with paler knob.
Legs: Yellow and pale brown setulose, hind trochanter normal, claws and pulvilli normal, mid tibia with strong ventral seta at apex. I cannot make out any row of short spines on the top of the front femora as described by Lamb (1914).
Abdomen: tg dark brown, dull with brown dusting, tg 5 with dorsal elongated quadrate brown spots, otherwise yellow; setulae brown to black about half length of tg. Postabdomen: tg 6 narrow and brown, cerc very small, narrow and pale, pregenital st broad with deep excavation fringed with dense long pale setulae, surs visible in this space as 2 short black sinuous projections.
As male, but tg 6-8 each with 2 brown almost triangular spots narrowly separated at middle. Postabdomen: tg 7 complete and st 7 thinly sclerotized; tg 8 in two small lateral sclerites; st 8 larger than tg 8 and microtrichose; hypr distinct, trapezoidal and of similar size to small cerc, both structures with fine setulae.
Length: [male] and [female] body 1.5 mm, wing 1.5 mm.
Type material examined: SEYCHELLES: [male] (holotype) "Mahe 08-9, Seychelles Exp. / Prof. J.S. Gardiner 1914-537 / A. sexvittata Lamb [handwritten] / Type H.T." [on a red bordered label] (BMNH); 3 [female] (paratypes) same data (19 BMNH, 29 CUMZ). Note: The holotype is in a reasonable condition: the pin damages only a part of the right half of the thorax. It has "spec 120" written on the mount.
Other material examined: MAURITIUS: 1 [male] Wolmar, 1-2 km S, coast roadside, 1.vi.2000, J.W. Ismay (OXUM). SEYCHELLES: 1 [female] Aldabra, West I., at light, nr settlement, 30-31.i.1968, B. Cogan & A. Hutson (NHML); 3 [male] 3 [female] Aldabra, West I., at light, nr settlement, 21-31.iii.1968, B. Cogan & A. Hutson (NHML); 19 Seychelles, North I., Calophyllum (Clusiaceae) woodland, 30.vii.2000, Malaise trap, J. Gerlach (NMWC).
Distribution: Mauritius, Seychelles.
Genus Tethysimyia gen. n.
Type species: Aphaniosoma deemingi Ebejer, 1996, here designated.
Etymology: From the Greek name Tethys (the sea goddess in mythology) and myia (fly), denoting the affinity of this genus with seashore habitats.
Diagnosis: Very small pale flies with rather large round head or head higher than long, very broad gena, well-developed, but narrow facial carina not reaching buccal margin, thus lower part of face between anterior eye margins flat, more or less square and carina therefore less conspicuous than in Krifomyia; recessed antenna, but not in deep fossa; chaetotaxy generally reduced and fine, 2 very short orb set high on fr, 1 dc, scutal setulae in 4-10 rows between dc lines, absent pra and sa setae, but post ia, pa and katepisternal present even though short; [R.sub.4+5] and [M.sub.1+2] slightly convergent; male postabdomen exceptionally small with dorsally narrow, complete epandrium; female with poorly sclerotized s, pigmented or unpigmented.
Head: More or less round and rather large in proportion to thorax, fr broad and moderately convex and protruding in front of eye margin, eye round or only slightly oval, gena broad, at least half height of eye, finely setulose, ocp concave though not quite as much as in Aphaniosoma, face poorly sclerotized with distinct carina in upper half between antennae, antennae small with third segment rounded, arista 2-segmented, about twice as long as rest of antenna, thickened at base, but rapidly becoming thinner, mouthparts small, palp minute oval and finely setulose, labella not elongated, setulae along buccal margin, including those at vibrissal corner, short and fine. Chaetotaxy: 0-3 very short orb, 1 vti and 1 vte both short, pvt minute or absent.
Thorax: Rather rounded, not elongated, scut of normal shape. Chaetotaxy: 1 pprn, 1 posthu, ihu present or absent, 2 ntpl, 1 dc, scutal setulae very fine and pale in 4-8 rows between dc lines, pra and sa absent, 1 post ia, 1 pa, 1 anepisternal, 1 katepisternal, anepisternal and katepisternal setulae scanty and extremely fine or minute, scut with usual 4 marginals.
Wing: Hyaline, relatively short and broad, veins very pale yellow, hu break present, subcosta merges with [R.sub.1] before latter joins costa, no erect spinose setulae on dorsal aspect of costa, [R.sub.4+5] and [M.sub.1+2] slightly convergent towards apex. Haltere short, pale with pale knob.
Legs: Slender, femora not dilated, segments of all legs pale and finely setulose, mid tibia with apicoventral seta longer than diameter of tibia at apex, hind metatarsus sometimes dilated, hind trochanter normal.
Abdomen: More or less oval and slightly dorsoventrally compressed, 6 segments in male and 7 in female, tg with fine setulae, st poorly sclerotized.
Postabdomen: Atypical for Aphaniosominae and closer to that of Krifomyia, with reduced modification of hypopygial structures, which are simple and partially within small ep; ep distinct from tg 6 and surs fused to it, apparently formed of extension of inferior margin.
Distribution: Coasts and islands of the Gulf of Mexico, Red Sea (Fig. 70), Indian and Pacific oceans.
Ecology: Probably associated with saline marshes and mangroves in coastal localities, in hot, tropical and subtropical climates.
Tethysimyia deemingi (Ebejer, 1996), comb. n. Figs 60, 61
Aphaniosoma deemingi: Ebejer 1996: 289.
This small species was described from a female taken in Oman. With additional specimens of both sexes now available, its distinctive characters may be better appreciated. The male is described and the postabdomen of both sexes is illustrated.
Head: Yellow, large and higher than long, fr broad with barely converging eye margins, antenna small and recessed, gena with numerous setulae and about as broad as height of eye, which is round, numerous fr setulae of equal length, 1 orb set far back and hardly longer than fr setulae, pvt minute.
Thorax: Yellow with faint deeper yellow longitudinal stripes on scutum. Chaetotaxy: numerous fine setulae over whole dorsal surface, 1 pprn, 1 short posthu which is a little less than twice length of adjacent setulae, 1 long dc and a shorter one in front, 1 short prscut pair of acrs, others not differentiated from general scutal setulae, scut with usual 4 marginals,prpl setula absent (specimen pinned through sides, but in other specimens anepisternal seta present and katepisternal absent).
[FIGURE 60 OMITTED]
[FIGURE 61 OMITTED]
Wing: Translucent and slightly iridescent with broad anal margin; veins very pale yellow to white; [R.sub.2+3] and [R.sub.4+5] run more or less parallel to each other and distance between them on costa about 0.9 that between [R.sub.4+5] and [M.sub.1+2], which converge towards apex; costa with stouter spine-like setulae and very few thin setulae; costa thinned out but not broken just beyond hu crossvein, subcosta vestigial, visible only as a fold. Haltere pale yellow.
Legs: Not dilated or modified, entirely pale yellow and short yellow setulose; tibia 2 with dark apicoventral seta.
Abdomen: Yellow with light brown bands on middle part of tg 2-5; very short setae on disc of tg, longer and stronger on tg 6; st largely membranous.
Postabdomen: Very small and inconspicuous in dry specimens; tg 6 longer than tg 5 when viewed from above, ep narrow dorsally, but broad laterally and ventrally such that most hypopygial structures lie within its boundaries; cerc relatively large and sclerotized with numerous and very fine setulae, hyp forming bridge between lower incurved margins of ep, surs in form of short lobe arising from posteromedial border of ep, prg very small, arising from posterior edge of hyp, psg not identified, ph simple with short narrow ph apd.
Similar to male in external characters. Postabdomen: tg 7 complete, tg 8 small and with minute setulae along anterior border; st 8 larger and with 2 setae at posterior border, ventrally; hypr distinct with fine setulae, cerc small and round; s poorly sclerotized, conical and with first part of duct thinly sclerotized.
Holotype (examined): OMAN: [female] Dhofar, Khor Rouri [Khawr Rawri, 17[degrees]03'S:54[degrees]26'E], 12.xi.1992, J.C. Deeming (NMWC).
Other material examined: EGYPT: 1 [male] Sinai, Taba, 1.v.1996, A. Freidberg (TAUI); 3 [female] Sinai, Taba, 10 km S, 1.v.1996, I. Yarom (TAUI); 1 9 Sinai, Ras Umm Burqa', 1.v.1996, I. Yarom (TAUI). OMAN: 1 [female] N Masirah Island, BERS Camp, 5-7.iii.1995, S.P. Dance (NMWC).
Distribution: Egypt, Oman.
The systematic position of the genera within the Chyromyidae is explored further with a cladistic analysis taking the Heleomyzinae as the outgroup taxon. D.K. McAlpine's proposal (McAlpine 2007) to categorize the Sphaeroceridae and Heleomyzidae into one family, the Hetermyzidae, and consequently alter their rank to subfamily, is not relevant to this analysis, for which I accepted the categorization of the Heleomyzinae as a subfamily. The Heteromyzidae (= Heleomyzidae and Sphaeroceridae) and the Chyromyidae share synapomorphic characters that justify retaining the latter family in closer phylogenetic proximity to the former rather than to any other acalyptrate family, but there are also characters that the Chyromyidae share with Ephydroidea and Opomyzoidea. However, the last two superfamilies have evolved autapomorphies (for example, several antennal characters, one proclinate fr orb seta, precoxal bridge present, short rigid ph) that distance them from the Chyromyidae. The few similarities have probably co-evolved, for example, spinose costa, absent pterostigma and reduction in spermathecae from three. At present, I find no reason to differ from the opinions of previous authors (D.K. McAlpine 1985; J.F. McAlpine 1989) that the Chyromyidae are closer to the Heteromyzidae.
This study of the African Chyromyidae uncovers more diversity than was hitherto appreciated, although many of the characters are not immediately obvious, particularly if low magnification microscopy is used. Some of these characters are present in the Heteromyzidae, though not uniformly so, and they may not all be homologous. For example, an incurved seta placed anteriorly on the fr vitta and not on the orb plate is found in Elachisoma Rondani, 1880 (Sphaerocerinae, Limosinini) and Tapeigaster Macquart, 1847 (Heteromyzidae, Tapeigastrinae), but this is not homologous with the incurved seta of Chyromyinae, which is inserted on the orb plate and therefore is a true orb. An anterior incurved seta on the scutum is present in Thoracochaeta Duda, 1918 (Sphaerocerinae, Limosinini) and in the Aphaniosominae, though it probably is not a homologous structure. I have called this the ihu seta and not a dc seta because it is not at all in line with the dc row. The anepisternal seta is always present in the Chyromyidae and it is found in Fenwickia Malloch, 1930 and Waterhouseia Malloch, 1936 (Heleomyzinae). The modified setulae of the anterodorsal aspect of the costa, found in many Heleomyzinae, are commonly also found in some Chyromyinae and rarely in Sphaerocerinae (Elachisoma). The costal break near the hu crossvein, though very rare in Heleomyzinae, is present in a number of species, e.g., Diplogeomyza spinosa McAlpine, 1967 (Australia), Blaesochaetophora picticornis Bigot, 1888 (Chile) and Gephromyza sp. (Chile). It is present in many Sphaerocerinae, and within the Chyromyidae, it is universally found in Aphaniosominae. Marshall and Richards (1987) state that the wing in Sphaerocerinae has costagial, hu and subcostal breaks. They illustrate examples. Rohacek (1998) repeats this statement and gives the same illustrations. I examined a number of species in a range of genera from all over the world and I find the hu vein is an unstable character (indeed, also so in the Chyromyidae), there being genera that include species with no break or with a complete break. Often there is a partial break, i.e., a narrowing and/or desclerotization, but no actual interruption. For example, Lotobia moyoensis Vanschuytbroek, 1959 (Nigeria) has a complete break, L. elegans Vanschuytbroek, 1959 (Nigeria), a partial break, L. elegans Kim & Han, 1990 (Nigeria), no break. A break at the hu crossvein is absent also in Copromyza fumipennis Stenhammar, 1854 (Pakistan), C. kibokoensis Vanschuytbroek, 1959 (Congo), C. marginatis Adams, 1905 (Oman), Crumomyia glabrifrons (Meigen, 1830) (France), and C. nitida (Meigen, 1830) (Sicily). A complete break is present in Pterogramma sp. (Malaysia) and Sphaerocera sp. (Yemen). A partial break is present in Poecilosomella angulata (Thomson, 1869) (Morocco).
I propose a closer affinity of the Chyromyidae with the Heteromyzidae than with the Ephydroidea. In this, I agree with J.F. McAlpine (1989). Since I found that many characters of the Chyromyidae are represented in one or other of the subfamilies of the Heteromyzidae, namely the Sphaerocerinae and Heleomyzinae, it was unclear to me which of these might be more closely related to the Chyromyidae. In a preliminary analysis, I used both as outgroup taxa and the Heleomyzinae consistently branched out closer to the Chyromyidae than did the Sphaerocerinae. The former was therefore chosen as the outgroup taxon for the cladistic analysis of the new subfamily and generic classification of the Chyromyidae proposed in this study.
The groundplan conditions of the Chyromyidae that I recognize are listed below:
(1) pale integument
(2) face desclerotized centrally below antennal foveae except for a linear vertical ridge
(3) true vibrissa absent
(4) pvt setae convergent
(5) two or more reclinate orb setae
(6) setulose fr
(7) second antennal segment not slit dorsally
(8) prosternum narrow with median groove
(9) one or more dc seta
(10) posthu seta well-developed
(11) two subequal pairs of scut setae
(12) one anepisternal and one katepisternal seta
(13) anepimeron and meron bare
(14) tibiae without preapical posterodorsal seta and without any developed setae along shaft
(15) metatarsus of anterior, middle and posterior legs of more or less equal length
(16) wing with costal break at [R.sub.1]
(17) R1 not dilated at junction with costa
(18) subcosta merging with [R.sub.1] close to junction with costa
(19) veins [R.sub.2+3] and R convergent
(20) anal and posterior basal cells closed
(21) male abdomen with six and female abdomen with seven pregenital segments
(22) postabdomen symmetrical in both sexes
(23) female postabdominal segments barely extensile, st 8 divided and epiproct desclerotized
(24) two spermathecae
(25) male hypopygium with U-shaped hyp
(26) surs, prg and psg not fused with their respective basal hypopygial sclerites
The autapomorphies that characterize the Chyromyidae with respect to the Heteromyzidae are: a pale yellow integument; membranous face; vibrissa absent; wing with veins [R.sub.2+3] and [R.sub.4+5] convergent; dorsal preapical tibial seta absent; postabdomen symmetrical in both sexes; two spermathecae; articulated surs; prg and psg; and halophilous habits. Synapomorphies shared with the Heteromyzidae include: discoid third antennal segment; second antennal segment without dorsal slit; orb setae reduced to 2 or 3; subcosta reaching costa or merging with [R.sub.1] at junction with costa; subcostal break present; [A.sub.1] not reaching wing margin; distiph complex and enlarged; female postabdomen not or barely extensile.
The much greater diversity within the Heteromyzidae adds to the problem of interpreting the polarity of characters, although ultimately this would not affect the result of the cladistic analysis. For the purpose of this exercise, I have limited myself to relatively few and easily interpreted characters. I examined a limited number of species from a range of genera of Heleomyzinae and Sphaerocerinae from across the world, but for the Chyromyidae, I examined all the genera and, within each, a great majority of the species (several of them undescribed) from most zoogeographical regions (Table 2). Where I was unable to examine specimens, I did not rely on descriptions, as some of the chaetotaxic characters were either not mentioned or are subject to misinterpretation or at least to an interpretation different from mine.
The only genus, indeed the only species, of the Chyromyidae ever properly considered for phylogenetic study has been Chyromya flava. The other genera have not been considered in any detail and, with regard to the male and female postabdominal characters, have not been considered at all. At the very least, Aphaniosoma must now be taken into account as this significantly widens the concept of the Chyromyidae. Aphaniosoma B (Table 2) consists of those species with a strong presutural dorsocentral seta and other defining features that possibly justify raising this group to generic status. The cladistic analysis reported in this article lends support to this. However, this requires further investigation and is beyond the scope of the present study, as several more species awaiting description are known to me.
For each character assessed within the Chyromyidae, I accepted only 100 % concordance among all the species examined within each genus, with one exception. In Gymnochiromyia, a presutural dc seta is found in a minority of species. This being the only character state that I found to be at odds with the remaining characters of the genus, I considered it a reversal when present. The interpretations of the character states are listed below. The numbering starts at zero to match the first character as it appears on the cladogram generated by the program TNT. Cladistic analysis was performed using TNT version 1.1 (Goloboff et al. 2008). This program is made available for use by the Willi Hennig Society (http://www.zmuc.dk/public/phylogeny/TNT; downloaded 15.iii.2008).
Character 0. Concave ocp. Across the order, and particularly in the Heleomyzinae, a convex ocp is the usual form. Therefore, I consider a concave ocp an apomorphy within the Chyromyidae.
Character 1. Postgenal seta. This small seta or setula lies posteroventrally on the inferior aspect of the ocp region of the head. It is a subtle character in the Chyromyidae because of their small size. It is rarely mentioned in descriptions of other acalyptrates. Nevertheless, it is widespread and common in many families. I consider its loss in some Chyromyidae an apomorphy.
Character 2. Incurved lower fr-orb. A very few Sphaerocerinae exhibit this character state (seta on the fr vitta), but I consider it a homoplasy rather than a homology with the character state (seta on the orb plate) in the Chyromyinae (in which taxon it is universal). I consider it an apomorphy in the Chyromyidae, it being even more unusual in the other acalyptrates.
Character 3. orb reduced to setulae or absent. Having well-developed orb is the usual character state in the great majority of acalyptrates. Wherever a loss or reduction occurs, often with loss or reduction of other somatic setae, it is considered to be an apomorphic state.
Character 4. Pair of long fr setae. This pair of usually divergent setae is placed in the middle of the fr in front of the oc triangle. It appears sporadically in a number of genera in several families of the acalyptrates. I consider it an apomorphy within the Chyromyidae as it appears in only one species group.
Character 5. Disc of ocp bare. Here, I refer to the space on the ocp between the postocular row(s) of setae or setulae and those situated strictly along the margin of the ocp foramen. This character is common in the Heleomyzinae and Sphaerocerinae, as well as in many other acalyptrates. I therefore consider this absence of setae within the Chyromyidae an apomorphy.
Character 6. pvt setae reduced to setulae or absent. Across the whole suborder, the usual character state is the presence of pvt setae. Their absence, being also uncommon within the Chyromyidae, confirms their apomorphic state.
Character 7. Well-developed facial carina. Although this character state appears in a number of genera within a range of families of the acalyptrates, it is not the norm. A tendency to form a shallow carina is common within the Chyromyidae, but its marked development in some cases suggests this is an apomorphy.
Character 8. Absent vibrissa. The loss of the vibrissa is uncommon in the Schizophora and in the Chyromyidae this is an apomorphy relative to the outgroup.
Character 9. Costa with a complete hu break. This character is often found in the Sphaerocerinae, but rarely in the Heleomyzinae. It is an apomorphic state in the Schizophora. Within the Chyromyidae, it occurs in the four genera of Aphaniosominae, but is rare and incomplete in the Chyromyinae. I consider it an apomorphy within the family.
Character 10. Spine-like setulae on costa. These are common in the Heleomyzinae and rare in the Sphaerocerinae. They are a feature of the Chyromyinae, but not the Aphaniosominae. I consider them to be synapomorphic for the Chyromyinae and Heleomyzinae.
Character 11. [R.sub.2+3] and [R.sub.4+5] convergent. The convergence of these two longitudinal veins is an apomorphy.
Character 12. [R.sub.4+5] and [M.sub.1+2] divergent. Although this is the norm in acalyptrates, I consider it to be possibly an apomorphy within the Chyromyidae (a reversal?).
Character 13. Absent presutural dc seta. This is an apomorphy among the higher Diptera. In Gymnochiromyia it is most often absent and I consider its presence in the minority of species a reversal.
Character 14. prsut ia seta. This is not the same seta referred to by Rohacek (1998) as synonymous with the posthu seta in the Sphaerocerinae. In the Chyromyidae, all species have a true posthu. The true prsut ia, when it occurs, is inserted medially and slightly posterior to the posthu. This is a reversal among the higher Diptera especially among the acalyptrates. Therefore, in the Chyromyidae it is likely an apomorphy.
Character 15. Lost or reduced post ia seta. A post ia seta is present in most Sphaerocerinae and Heleomyzinae. Its loss in the Chyromyidae is an apomorphy.
Character 16. Absent postsutural pra seta. This seta is very common in many higher Diptera. I consider its loss in most Chyromyidae an apomorphy, but equally it could be considered a reversal, given its presence in only two genera.
Character 17. Absent sa seta. A seta in this position is a very widespread character in the Diptera. I consider its loss within the Chyromyidae an apomorphy.
Character 18. scut setae reduced to 2 pairs of marginals is the plesiomorphic state in the acalyptrates. The increase in number in a few genera of the Chyromyinae, and also in a few genera of Heleomyzinae and Sphaerocerinae, suggests that the presence of more than two pairs is apomorphic.
Character 19. Absent setulae on disc of scut. This is probably an apomorphy among the higher Diptera and within the Chyromyidae. These setulae are not of the same structure as very short setae.
Character 20. acrs setulae in 2 rows. While there is a very wide range of states among the acalyptrates, from a completely bare to a densely setulose scutum, at least in comparison to the outgroup taxa, reduction to two rows of setulae appears to be an apomorphy.
Character 21. Absent prpl seta/setula. An apomorphic state among the acalyptrates.
Character 22. Absent anepisternal seta. An apomorphic state among the acalyptrates; absent in most genera of the Heleomyzinae and in almost all Sphaerocerinae.
Character 23. Absent katepisternal seta. This seta is most often present in higher Diptera. I consider its loss among the acalyptrates and within the Chyromyidae an apomorphic state.
Character 24. Dilated femora. This feature occurs sporadically across a wide range of acalyptrates. I consider it an apomorphic state among the Chyromyidae.
Character 25. Mid tibia with reduced or absent apicoventral seta. A majority of Diptera in all families have one or more such setae. Their absence is an apomorphy.
Character 26. Tibia without dorsal preapical seta. The presence of this seta is generally considered to be an apomorphy among the Heteromyzidae. Its loss in the Chyromyidae is probably secondary (a reversal) and therefore an apomorphy relative to the chosen outgroups.
Character 27. Internalised ep. This highly complex development in Aphaniosoma is undoubtedly an apomorphic state.
Character 28. Modified cerc in males. This is an apomorphic state in acalyptrates and the Chyromyidae.
Character 29. Loss of ej apd. I consider this an apomorphic state within the Chyromyidae, given its presence in one subfamily and in the chosen outgroup taxa.
Character 30. ph apd extensively fused with hyp. The plesiomorphic state expecially in the outgoup taxon is a long and narrow ph apd that remains free from the hyp almost to its apex, where it articulates with the basiph.
Character 31. Two spermathecae. This is an apomorphic state in the Chyromyidae.
Character 32. Female postabdomen with well-developed fleshy lobes medial to, and attached to, sclerites of st 8. This subtle character is found in only one genus of Chyromyinae (Somatiosoma). I consider it an apomorphy.
The characters listed above are given in the matrix (Table 2) used for the analysis. Parsimony analysis using "Implicit enumeration" methodology in TNT generated three equally parsimonious trees (score 45) of identical topology with a different interpretation of only a small number of characters. The only "phylogenetic" difference between them was the placement of the genera Oroschyromya, Notiochyromya and Somatiosoma relative to each other. The strict consensus tree is depicted in Fig. 62. Tree 0 was one step shorter than the others and it was supported by the cladogram generated by the Majority Rule (Fig. 63). It was therefore selected and the synapomorphies were superimposed on the clades (Fig. 64). Bootstrap support was calculated using 1,000 replicate routines and the results superimposed on the strict consensus tree (Fig. 65). Manual calculations on tree 0 gave a length = 28; a consistency index, CI=0.893; a retention index, R=0.942; and a rescaled consistency index, RC=0.841.
This analysis provides support for the proposed generic grouping into two subfamilies and gives an indication of the phylogenetic relationships of the genera of the Chyromyidae. The tree supports well the phylogenetic position of the genera within the Aphaniosominae, with bootstrap scores of 87 % or above, with the exception of Paraphaniosoma. I consider their relationship to each other, as well as to the subfamilies, to be well resolved. Paraphaniosoma presents at least two very strong characters that convince me it belongs to the Aphaniosominae close to Aphaniosoma A and Aphaniosoma B, rather than Tethysimyia or Krifomyia. These characters are the arrangement of structures of the hypopygium and those of the head. The presence of the pra seta is most probably a reversal or homoplasy. Support is weak for the genera Gymnochiromyia, Oroschyromya, Notiochyromya and Somatiosoma. On general appearance and detailed examination of the postabdomen of both sexes, it appears to me that Notiochyromya is closest to Chyromya, and Oroschyromya closest to Gymnochiromyia. The position of Somatiosoma is as yet unclear, although I suspect it will eventually turn out to be an offshoot of Notiochyromya. At this stage, I cannot consider that the relationship of these four genera to each other or within the Chyromyinae is resolved. This may be partly due to not using more characters of the male and female postabdomen and to an avoidance of other somatic characters where definition is difficult, for example, the relative head shape: prognathous versus opisthognathous, the degree of convergence of eye margins, and the subtle wing characters. These three genera are actually quite easy to separate (with a little practice and experience) in both sexes without dissection, relying only on somatic as opposed to hypopygial structures. To resolve the phylogenetic relationships within the Chyromyinae, a study of more species from other regions of the world may be helpful, but most of the species are undescribed. The choice of an outgroup taxon to perform such an analysis is not easy, as some somatic and hypopygial structures, though clearly very differently developed in the Aphaniosominae, are not easily defined in a categorical way across the family (i.e., present or absent) as is required to run a cladistic analysis. For example, the breadth and depth of the basal half of the phallapodeme is developed differently in the Chyromyinae. It may also be described as broad and deep in Aphaniosoma, but here it clearly presents a very different modification. The same applies to the modification of the ejaculatory apodeme in the genera of the Chyromyinae.
[FIGURE 62-63 OMITTED]
[FIGURE 64-65 OMITTED]
Support for the monophyly of the Chyromyidae is very good, with a supporting bootstrap value of 100%. Similarly, the division into two subfamilies appears to be well supported, with a bootstrap value of 93%. Based on the chosen characters and the interpretation of their polarity, the Chyromyidae appear to be closer to the Heleomyzinae rather than the Sphaerocerinae. However, a further cladistic analysis using a larger character set, may be needed to establish the relationships between these and other closely related taxa, such as the Ephydroidea.
ZOOGEOGRAPHY AND ECOLOGY
The family is known from all continents and zoogeographical regions except the Arctic and Antarctica (Cogan 1980; McAlpine 1965; Pitkin 1989; Soos 1984; Steyskal 1977). I have seen scanty material from mainland South America, but I know of much more collected material that awaits study. I am unaware of any published information other than a study on the Galapagos species (Wheeler & Sinclair 1994). Specimens have been collected on a number of remote Atlantic and Pacific islands (Frey 1945; Hardy & Delfinado 1980). Many are still undescribed and therefore zoogeographical affinities are speculative at best. The distribution of the genera of Chyromyidae in Africa is summarised in the maps given in Figs 66-70.
[FIGURE 66 OMITTED]
[FIGURE 67 OMITTED]
The genus Aphaniosoma is widespread in the Holarctic and Afrotropical regions, with most species diversity being in the Mediterranean (Carles-Tolra 2001; Ebejer 1998; 2005; Ebejer & Baez 2001), Central Asia (Ebejer 2006) and southern Africa (this paper), in that order. There are some affinities between groups of species in the eastern Mediterranean and Africa and between other groups of species in Central Asia and the Mediterranean, but not between Africa and Asia. On current knowledge, therefore, it appears that perhaps most species radiation has occurred from the northern part of Africa (e.g., A. approximatum and A. gallagheri). The central Sahara and the tropical rainforests of central Africa may have been early separating factors for some species groups, whereas the Rift Valley and the savannah zone from southwest Africa to northeast Africa may still act as natural corridors facilitating the spread of species (e.g. Aphaniosoma fissum, Somatiosoma eremicolum, S. nitescens and Oroschyromya affinis). The countries between the Mediterranean part of the Middle East and the central Asian states of Mongolia and the former Soviet Union have not been investigated. For example, Iraq, Iran, Afghanistan and Pakistan may be rich in species of this genus.
The genera Somatiosoma and Oroschyromya appear to be typically Afrotropical, but the Indian subcontinent has not been investigated for Chyromyidae. If they were truly restricted to the Old World tropical and subtropical areas, both genera might still be expected to occur in the Indian subcontinent. Somatiosoma favours hot, arid habitats and such habitats occur in northwest and central India. The genus has been found in all parts of the Arabian Peninsula. Oroschyromya, with its apparent affinity for wetter regions, might occur in the Western Ghats. Notiochyromya, on the other hand, is represented in all geographical areas of the southern hemisphere and as studies continue and more species are discovered, this genus may turn out to be rich, diverse and inhabiting a wide range of habitats extending into Central America and the southern states of the USA. The Chyromyidae of the Oriental and Australasian regions (Steyskal 1977; Pitkin 1989) are too poorly known for further comments to be made at present.
[FIGURE 68 OMITTED]
[FIGURE 69 OMITTED]
[FIGURE 70 OMITTED]
As far as is known, most speciation in Gymnochiromyia appears to be in the Mediterranean (Ebejer 1998) and in southern Africa (Ebejer 2008b). This suggests a disjunction and lends support to the theory that the African fauna includes the Mediterranean, with the Sahara being only a recent barrier (Kirk-Spriggs & McGregor 2009). The genus, as originally interpreted, has been recorded from North America. I have seen limited material. Very little work has been done on Nearctic Chyromyidae (McAlpine 1965, 1987) and it is difficult to speculate as to the richness or otherwise of the fauna there, or even as to what affinities this might have with the fauna of Europe and Africa.
This paper gives descriptions of 32 new taxa consisting of five new genera and 27 new species. Thus, the total number of species of Chyromyidae described worldwide is now almost 180.
The African species of Chyromyidae provide further evidence of the remarkable diversity within the family and may offer a better understanding for the generic classification of species. This diversity provides a platform for a more detailed study of the large genus Aphaniosoma, wherein many more species known to the author await description and more species groups are apparent than has been given in the preliminary cladisitc analysis above. However, it is not yet clear how these species groups relate to each other and whether or not there will be significant grounds for further taxonomic subdivision.
Although it was not practicable to examine every known species worldwide for this study, many from all zoogeographical regions were examined, and the generic limits as defined in this article apply to all the specimens that I have seen.
In general, Chyromyidae are most speciose, and populations are largest, in lowlands. The Afrotropical species occurring at a high altitude similar to that where Aphaniosoma species occur in central Asia (Ebejer 2006) belong to the new genera in the Chyromyinae and not to the Aphaniosominae. This is interesting in as much as it may suggest evolutionary divergence in the subfamily rather than these being relict species or species groups that became isolated in the African highlands. This is supported by the structural modifications in the Afrotropical species (e.g., species in the genera Oroschyromya and Notiochyromya), that have no counterparts in species from elsewhere. Species of Aphaniosoma from the Mediterranean and the Middle East appear to share characteristics with the Afrotropical species. A few species have distributions that extend from southern Africa to East Africa and to Arabia, supporting the view that dispersal has occurred along the Rift Valley and across the savannahs of Africa (Kirk-Spriggs & Stuckenberg 2009). However, little more can be deduced until the countries encompassing the Sahara are investigated. More isolated mountainous areas need to be sampled to ascertain the prevalence or otherwise of relict species. This is especially so for the Haggar mountains in Algeria, Tibesti in Chad, Darfur in Sudan and the mountains of Ethiopia.
This study was based on material from only 24 African countries, many of which have not been adequately sampled for Chyromyidae. Thus, most of the continent remains effectively unexplored. Some species are recorded from localities very far apart across the continent, suggesting that they might easily occur in the intervening regions where similar habitats occur. For example, species such as Aphaniosoma aethiops, A. approximatum, A. fissum, A. gallagheri, Notiochyromya filigera, N. sexspinosa, Oroschyromya affinis, Somatiosoma eremicolum, S. grandicornis and S. nitescens are likely to be much more widspread than records in this article suggest, and it is already apparent that S. nitescens probably occurs all over Africa.
I am very grateful to the many colleagues and authorities of institutions who kindly made available to me the specimens in their care: David Barraclough, Durban, South Africa; Roy Danielsson, Lund, Sweden; John Deeming, Cardiff, UK; William Foster, Cambridge, UK; Amnon Freidberg, Tel Aviv, Israel; John Ismay, Oxford, UK; Ashley Kirk-Spriggs, Bloemfontein, South Africa (formerly of Windhoek, Namibia); Marion Kotrba, Munich, Germany; Darren Mann, Oxford, UK; Eugene Marais, Windhoek, Namibia; Rudolph Meier, Singapore (formerly of Copenhagen, Denmark); Bernhard Merz, Geneva, Switzerland; Mike Mostovski, Pietermaritzburg, South Africa; Thomas Pape, Copenhagen, Denmark; Adrian Plant, Cardiff, UK; Michael von Tschirnhaus, Bielefeld, Germany; Pekka Vilkamaa, Helsinki, Finland; Nigel Wyatt, London, UK; Joachim Ziegler, Berlin, Germany. James Turner (Cardiff) gave much appreciated practical advice and assistance with the preparation of the photomicrographs of the wings. A. Plant was most helpful with general discussion on cladistics and specifically with running and interpreting TNT to produce meaningful phylogenetic trees. I am especially indebted to the following for their patient and critical review of earlier drafts of the manuscript: J. Deeming, Paul Gatt (Malta), and A. Kirk-Spriggs. A. Kirk-Spriggs exhibited untold patience checking the data and offered constructive comments. Three reviewers meticulously picked their way through the manuscript, identified errors and made many useful suggestions. The manuscript has greatly improved as a result, and I am most grateful for their efforts.
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Wheeler, T.A. & Sinclair, B.J. 1994. Chyromyidae (Diptera) from the Galapagos Islands, Ecuador: three new species of Aphaniosoma Becker. Proceedings of the Entomological Society of Washington 96 (3): 440-453.
Martin J. Ebejer Research Associate, Entomology Section, Department of Biodiversity & Systematic Biology, National Museum Wales, Cathays Park, Cardiff, CF10 3NP, UK; email@example.com
TABLE 1 Comparison of the somatic, chaetotaxic and hypopygial characters of the two subfamilies. Chyromyinae Aphaniosominae Head Straight or convex; Concave whether viewed face always flat, never from above or from the with carina even side; face variable, between antennae; palp often with median short oval to round, carina, even if very appearing disc-shaped; low, sometimes strongly arista with 3 segments. developed; palp elongate oval, almost club-shaped; arista with 2 segments, except in Krifomyia. Always 3 long orb, with 0-6 orb, none anterior one inclinate inclinate, usually relative to two short and becoming posterior ones and all progressively shorter more or less equally anteriorly; when 2 orb spaced from each other present, these often on fronto-orbital long and inserted in plate. upper half of fronto- orbital plate. Thorax Numerous setulae on all Setulae sparse; almost parts of scutum. totally restricted to acrs, dc and ia lines in Aphaniosoma; more numerous in other genera. No inclinate seta at Inclinate ihu seta or ihu location (at at least a setula anterior end of scutum present; reduced between pprn and most (apparently anterior dc). secondarily) in Krifo- myia and Tethysimyia. Strong anepisternal Anepisternal seta seta placed at middle placed in upper 1/3 of of hind margin of hind margin of anepisternum. anepisternum. No prpl seta or setula. prpl setula (rarely a seta) present, often extremely short. Legs One or more pairs of Femora never swollen; femora swollen to a apicoventral seta on greater or lesser mid tibia always extent (except in Oros- present and longer than chyromya); apicoventral diameter of tibia at seta on mid tibia, if apex. present (rarely), not longer than diameter of tibia at apex. Wing Veins always paler Veins darker towards towards posterior half apex of wing and vein of wing; vein separating basal and separating basal and discal cells always discal cells same palest; costa never colour as adjacent with spinose setulae on veins; costa often with dorsal aspect. spinose setulae on dorsal aspect. Male tg 1-6 always of normal tg 6 very often abdomen development and noticeably modified appearance, even if tg with secondary sexual 6 somewhat shorter and characters; sometimes often partly hidden also tg 4 and/or 5 below tg 5. modified. Male A more or less ep often divided into postabdomen spherical ep always right and left halves; present, distinct from almost always poorly and lying outside of tg visible because it lies 6. partly or wholly within tg 6; sometimes, viewed from behind (or from below), the two halves appear as shiny brown spots lateral to cerci. Cerci placed at lower Cerci placed at dorsal ventral aspect of ep aspect close to margin well away from dorsal of tg 6. edge of tg 6. hyp in profile hyp more variable, and generally broad (high), in most cases narrow in especially at base. profile. prg almost always a prg extremely variable, thin sclerite, setulose usually setulose on its external apically and typically (ventrolateral) attached to middle surface, attached to section of hyp after posterior lateral arm this articulates or of hyp before this fuses with ep, thus prg articulates with ep; lies medial and above thus it lies anterior inferior posterior and below inferior margin of ep. anterior angle of ep. psg usually small and psg extremely variable, loosely articulated closely and more firmly close to base of articulated to base of basiph, but rarely basiph, but often is so extends out of large as to extend well epandrial cavity; if it out of epandrial and does, then only tip sixth tergal cavity; in usually apparent. many Aphaniosoma clearly visible as black elongated structure. surs generally small, surs highly modified more or less hook- and often fused with shaped and loosely epandrium; may appear articulating with inner on any part of margin posteroventral margin of this sclerite. of ep. ph apd always long and ph apd usually broadly rod-shaped; basal half triangular with tbasal free with extreme apex part very short and somewhat trumpet- apex usually bi-lobed shaped and poorly (in dorsal view) and sclerotized. heavily sclero-tized. Separated ej apd Separated ej apd not present, variably apparent. sclero-tized. basiph short with a basiph very variable; sclerotized anterior when membranous, rodlike sclerite, downward and backward acting as fulcrum for extension of ph in most downward and backward species achieved extension of ph. through a turgid "inflation" and unfolding of whole ph; when sclerotized, mechanics similar to those in Chyromyinae. Female Fecund females very Fecund females without abdomen often with large ova such abdominal visible through distension and never integument of very with ova as clearly distended abdomen. visible as this. Female Spermathecae small and Spermathecae large, postabdomen round; tg 7 tending to variably shaped; tg 7 be divided, st 8 with 2 not tending to be distinct sclerites; divided and st 8 not hypr variable from well usually with 2 distinct to poorly sclero- sclerites; tg 8 often tized, rarely divided into small membranous. lateral plates that may lie anywhere from dorsal to ventral; hypr usually membranous. TABLE 2 Character matrix for the cladistic analysis of the genera of the Chyromyidae. Outgroup: Heleomyzinae. Abbreviations: char--character number, Sph--Sphaeroceridae, Hel--Heleomyzidae, Chy--Chyromya, Not -Notiochyromya, Oro--Oroschyromya, Som-- Somatiosoma, Gym--Gymnochiromyia, Aph A -Aphaniosoma A, Aph B- -Aphaniosoma B, Par--Paraphaniosoma, Tet--Tethysimyia, Kri-- Krifomyia. Character states: 0--plesiomorphic, 1--apomorphic; n--number of species examined (number known to the author in parenthesis). n 7(8) 12(14) 6(6) 7(7) char Sph Hel Chy Not Oro Som 0 0 0 0 0 0 0 1 0 0 0 0 0 0 2 0 0 1 1 1 1 3 0 0 0 0 0 0 4 0 0 0 0 0 0 5 0 0 0 1 1 1 6 0 0 0 0 0 0 7 0 0 0 0 0 0 8 0 0 1 1 1 1 9 0 0 0 0 0 0 10 0 1 1 1 1 1 11 0 0 1 1 1 1 12 1 1 0 0 0 0 13 0 0 0 1 0 1 14 1 0 0 0 0 0 15 0 0 1 1 0 1 16 1 1 1 0 1 1 17 0 0 0 0 0 0 18 1 1 1 1 0 0 19 0 0 0 1 1 0 20 0 0 0 0 0 0 21 0 0 1 1 1 1 22 1 1 0 0 0 0 23 0 0 0 0 0 0 24 0 0 1 0 0 0 25 0 0 1 1 1 1 26 0 0 1 1 1 1 27 0 0 0 0 0 0 28 0 0 0 0 1 0 29 0 0 0 0 0 0 30 0 0 0 0 0 0 31 0 0 1 1 1 1 32 0 0 0 0 0 1 23(25) 92(107) 14(16) 2(3) 9(9) 1(1) char Gym Aph A Aph B Par Tet Kri 0 1 1 1 1 1 1 1 0 1 1 1 1 1 2 1 0 0 0 0 0 3 0 0 0 0 1 1 4 0 1 0 0 0 0 5 1 1 1 1 1 1 6 0 0 0 1 1 1 7 0 0 0 0 1 1 8 1 1 1 1 1 1 9 0 1 1 1 1 1 10 1 0 0 0 0 0 11 1 1 1 1 1 1 12 0 1 1 0 0 0 13 1 1 0 1 1 1 14 0 0 1 0 0 0 15 1 1 1 0 0 0 16 1 1 1 0 1 1 17 0 1 1 0 1 1 18 0 0 0 0 0 0 19 1 1 1 1 1 1 20 0 1 1 0 0 0 21 1 0 0 1 1 1 22 0 0 0 0 0 0 23 0 0 0 0 0 1 24 0 0 0 0 0 0 25 1 0 0 0 0 0 26 1 1 1 1 1 1 27 0 1 1 1 1 1 28 0 0 0 0 0 0 29 0 1 1 1 1 1 30 0 1 1 1 0 0 31 1 1 1 1 1 1 32 0 0 0 0 0 0
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